{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"Q14676","secondaryAccessions":["A2AB04","A2BF04","A2RRA8","A7YY86","B0S8A2","Q0EFC2","Q2L6H7","Q2TAZ4","Q5JP55","Q5JP56","Q5ST83","Q68CQ3","Q86Z06","Q96QC2"],"uniProtkbId":"MDC1_HUMAN","entryAudit":{"firstPublicDate":"2005-07-05","lastAnnotationUpdateDate":"2026-01-28","lastSequenceUpdateDate":"2005-07-05","entryVersion":226,"sequenceVersion":3},"annotationScore":5.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"lineage":["Eukaryota","Metazoa","Chordata","Craniata","Vertebrata","Euteleostomi","Mammalia","Eutheria","Euarchontoglires","Primates","Haplorrhini","Catarrhini","Hominidae","Homo"]},"proteinExistence":"1: Evidence at protein level","proteinDescription":{"recommendedName":{"fullName":{"value":"Mediator of DNA damage checkpoint protein 1"}},"alternativeNames":[{"fullName":{"value":"Nuclear factor with BRCT domains 1"}}]},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"14695167"},{"evidenceCode":"ECO:0000312","source":"HGNC","id":"HGNC:21163"}],"value":"MDC1"},"synonyms":[{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"8724849"}],"value":"KIAA0170"},{"value":"NFBD1"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12475977"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12499369"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12551934"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607003"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607004"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607005"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12611903"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"14695167"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15201865"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15377652"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16049003"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16377563"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18411307"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18582474"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18583988"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18678890"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"30898438"}],"value":"Histone reader protein required for checkpoint-mediated cell cycle arrest in response to DNA damage within both the S phase and G2/M phases of the cell cycle (PubMed:12475977, PubMed:12499369, PubMed:12551934, PubMed:12607003, PubMed:12607004, PubMed:12607005, PubMed:12611903, PubMed:14695167, PubMed:15201865, PubMed:15377652, PubMed:16049003, PubMed:16377563, PubMed:30898438). Specifically recognizes and binds histone H2AX phosphorylated at 'Ser-139', a marker of DNA damage, serving as a scaffold for the recruitment of DNA repair and signal transduction proteins to discrete foci of DNA damage sites (PubMed:12607005, PubMed:15201865, PubMed:16049003, PubMed:16377563, PubMed:30898438). Also required for downstream events subsequent to the recruitment of these proteins (PubMed:12607005, PubMed:15201865, PubMed:16049003, PubMed:16377563, PubMed:18582474). These include phosphorylation and activation of the ATM, CHEK1 and CHEK2 kinases, and stabilization of TP53/p53 and apoptosis (PubMed:12499369, PubMed:12551934, PubMed:12607004). ATM and CHEK2 may also be activated independently by a parallel pathway mediated by TP53BP1 (PubMed:12499369, PubMed:12551934, PubMed:12607004). Required for chromosomal stability during mitosis by promoting recruitment of TOPBP1 to DNA double strand breaks (DSBs): TOPBP1 forms filamentous assemblies that bridge MDC1 and tether broken chromosomes during mitosis (PubMed:30898438). Required for the repair of DSBs via homologous recombination by promoting recruitment of NBN component of the MRN complex to DSBs (PubMed:18411307, PubMed:18582474, PubMed:18583988, PubMed:18678890)"}],"commentType":"FUNCTION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12551934"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607003"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607004"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607005"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12611903"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15201865"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15377652"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16049003"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16377563"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18006705"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18283122"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18411307"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18582474"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18583988"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18678890"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20008512"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20159462"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21482717"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22234877"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22635276"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23891287"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"30898438"}],"value":"Homodimer (PubMed:22234877). Interacts with H2AX, which requires phosphorylation of H2AX on 'Ser-139' (PubMed:15201865, PubMed:16049003, PubMed:16377563, PubMed:20159462). Interacts with the MRN complex, composed of MRE11, RAD50, and NBN (PubMed:12607003, PubMed:12607005). Interacts with CHEK2, which requires ATM-mediated phosphorylation of 'Thr-68' within the FHA domain of CHEK2 (PubMed:12551934, PubMed:12607004). Interacts constitutively with the BRCA1-BARD1 complex, SMC1A and TP53BP1 (PubMed:12611903, PubMed:22635276). Interacts with ATM and FANCD2, and these interactions are reduced upon DNA damage (PubMed:12607005). Also interacts with the PRKDC complex, composed of XRCC6/KU70, XRCC5/KU80 and PRKDC/XRCC7 (PubMed:15377652). This interaction may be required for PRKDC autophosphorylation, which is essential for DNA double strand break (DSB) repair (PubMed:15377652). When phosphorylated by ATM, interacts with RNF8 (via FHA domain) (PubMed:18006705). Interacts with CEP164 (PubMed:18283122). When phosphorylated, interacts with APTX (via FHA-like domain) (PubMed:20008512). Interacts (when phosphorylated) with TOPBP1; promoting TOPBP1 localization to DNA damage sites during mitosis (PubMed:21482717, PubMed:23891287, PubMed:30898438). Interacts (when phosphorylated) with NBN; promoting NBN and MRN complex localization to DNA damage sites (PubMed:18411307, PubMed:18582474, PubMed:18583988, PubMed:18678890)"}],"commentType":"SUBUNIT"},{"commentType":"INTERACTION","interactions":[{"interactantOne":{"uniProtKBAccession":"Q14676","intActId":"EBI-495644"},"interactantTwo":{"uniProtKBAccession":"Q13315","geneName":"ATM","intActId":"EBI-495465"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q14676","intActId":"EBI-495644"},"interactantTwo":{"uniProtKBAccession":"P38398","geneName":"BRCA1","intActId":"EBI-349905"},"numberOfExperiments":4,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q14676","intActId":"EBI-495644"},"interactantTwo":{"uniProtKBAccession":"P16104","geneName":"H2AX","intActId":"EBI-494830"},"numberOfExperiments":21,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q14676","intActId":"EBI-495644"},"interactantTwo":{"uniProtKBAccession":"O60934","geneName":"NBN","intActId":"EBI-494844"},"numberOfExperiments":33,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q14676","intActId":"EBI-495644"},"interactantTwo":{"uniProtKBAccession":"O96028","geneName":"NSD2","intActId":"EBI-2693298"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q14676","intActId":"EBI-495644"},"interactantTwo":{"uniProtKBAccession":"O96028-1","geneName":"NSD2","intActId":"EBI-15910280"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q14676","intActId":"EBI-495644"},"interactantTwo":{"uniProtKBAccession":"Q06609","geneName":"RAD51","intActId":"EBI-297202"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q14676","intActId":"EBI-495644"},"interactantTwo":{"uniProtKBAccession":"O76064","geneName":"RNF8","intActId":"EBI-373337"},"numberOfExperiments":11,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q14676","intActId":"EBI-495644"},"interactantTwo":{"uniProtKBAccession":"O76064-1","geneName":"RNF8","intActId":"EBI-15964690"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q14676","intActId":"EBI-495644"},"interactantTwo":{"uniProtKBAccession":"O43070","geneName":"nbs1","intActId":"EBI-2125045"},"numberOfExperiments":2,"organismDiffer":true}]},{"commentType":"SUBCELLULAR LOCATION","note":{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607005"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15201865"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16049003"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16377563"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20008512"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"30898438"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"35842428"}],"value":"Associated with chromatin (PubMed:12607005, PubMed:15201865, PubMed:16049003, PubMed:16377563). Relocalizes to discrete nuclear foci following DNA damage, this requires 'Ser-139' phosphorylation of H2AX (PubMed:12607005, PubMed:15201865, PubMed:16049003, PubMed:16377563, PubMed:30898438, PubMed:35842428). Colocalizes with APTX at sites of DNA double-strand breaks (PubMed:20008512)"}]},"subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12475977"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12499369"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12551934"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607003"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607004"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607005"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12611903"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"14695167"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15201865"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15377652"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20008512"}],"value":"Nucleus","id":"SL-0191"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607005"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15201865"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16049003"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16377563"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18411307"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18582474"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18583988"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18678890"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20008512"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21482717"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"30898438"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"35842428"}],"value":"Chromosome","id":"SL-0468"}}]},{"commentType":"ALTERNATIVE PRODUCTS","events":["Alternative splicing"],"isoforms":[{"name":{"value":"1"},"isoformIds":["Q14676-1"],"isoformSequenceStatus":"Displayed"},{"name":{"value":"2"},"isoformIds":["Q14676-2"],"sequenceIds":["VSP_014593"],"isoformSequenceStatus":"Described"},{"name":{"value":"3"},"isoformIds":["Q14676-3"],"sequenceIds":["VSP_034104"],"isoformSequenceStatus":"Described"},{"name":{"value":"4"},"isoformIds":["Q14676-4"],"sequenceIds":["VSP_014593","VSP_034103"],"isoformSequenceStatus":"Described"}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12499369"}],"value":"Highly expressed in testis"}],"commentType":"TISSUE SPECIFICITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12499369"}],"value":"Tandemly repeated BRCT domains are characteristic of proteins involved in DNA damage signaling. In MDC1, these repeats are required for localization to chromatin which flanks sites of DNA damage marked by 'Ser-139' phosphorylation of H2AX"}],"commentType":"DOMAIN"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607003"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607004"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12607005"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18411307"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18583988"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18678890"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22234877"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"30898438"}],"value":"Phosphorylated upon exposure to ionizing radiation (IR), ultraviolet radiation (UV), and hydroxyurea (HU) (PubMed:12607003, PubMed:12607004, PubMed:12607005). Phosphorylation in response to IR requires ATM, NBN, and possibly CHEK2 (PubMed:12607003, PubMed:12607004, PubMed:12607005). Also phosphorylated during the G2/M phase of the cell cycle and during activation of the mitotic spindle checkpoint (PubMed:12607003, PubMed:12607004, PubMed:12607005). Phosphorylation at Thr-4 by ATM stabilizes and enhances homodimerization via the FHA domain (PubMed:22234877). Phosphorylated at Ser-168 and Ser-196 by CK2 in response to DNA damage during mitosis, promoting interaction with TOPBP1 (PubMed:30898438). Phosphorylated by CK2 in response to DNA damage, promoting interaction with NBN and recruitment of the MRN complex to DNA damage sites (PubMed:18411307, PubMed:18583988, PubMed:18678890)"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22635276"}],"value":"Sumoylation at Lys-1840 by PIAS4 following DNA damage promotes ubiquitin-mediated degradation"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22635276"}],"value":"Ubiquitinated by RNF4, leading to proteasomal degradation; undergoes 'Lys-48'-linked polyubiquitination"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"24240613"}],"value":"Methylated at Lys-45. Demethylation at Lys-45 by JMJD1C after exposure to IR promotes MDC1-RNF8 interaction, RNF8-dependent MDC1 ubiquitination and recruitment of RAP80-BRCA1 to polyubiquitylated MDC1"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23891287"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"30898438"}],"value":"Phosphorylation at Ser-329 and Thr-331 by CK2 was initially thought to play a major role in the interaction with TOPBP1 (PubMed:23891287). However, it was later shown that phosphorylation at Ser-168 and Ser-196 promote interaction with TOPBP1 (PubMed:30898438)"}],"commentType":"CAUTION"},{"commentType":"SEQUENCE CAUTION","sequenceCautionType":"Erroneous initiation","sequence":"BAA11487.2","evidences":[{"evidenceCode":"ECO:0000305"}]},{"commentType":"SEQUENCE CAUTION","sequenceCautionType":"Erroneous termination","sequence":"CAH18685.1","note":"Truncated C-terminus.","evidences":[{"evidenceCode":"ECO:0000305"}]}],"features":[{"type":"Chain","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":2089,"modifier":"EXACT"}},"description":"Mediator of DNA damage checkpoint protein 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